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Journal of the Lepidopterists' Society
now its archaic condition threatens its survival. On the other hand Pieris napi has kept its evolutionary mobility and its subspecies have occupied a variety of environments, some still changing rapidly; in that sense the species has retained its youth.
Literature Cited
Bowden, S. R. 1964. The maintenance for experimental purposes of form "sul-phurea" of Pieris napi. J. Lepid. Soc. 18: 91-100.
Descimon, H. 1966. A propos de la plante nourriciere de Pieris ergane Geyer. Alexanor 4: 207.
Dethier, V. G. 1954. Evolution of feeding preferences in phytophagous insects. Evolution 8: 32-54.
Ehrlich, P. R. and P. H. Raven. 1964. Butterflies and plants: a study in co-evolution. Evolution 18: 586—608.
Hovanitz, W. and V. C. S. Chang. 1962. Three factors affecting larval choice of food plant. J. Res. Lepid. 1: 51-61.
Lorkovic, Z. 1941. Die Chromosomenzahlen in der Spermatogenese der Tag-falter. Chromosoma 2: 155—191.
----------■ 1968. Systematisch-genetische und okologische Besonderheiten von Pieris
ergane. Mitt, schweiz. ent. Ges. 41: 233-244.
Rothke, M. 1931. Einige Notizen uber Vorkommen und Lebensgewohnheit von Pieris napi L. in Nordamerika. Int. ent. Z. 25: 262^263.
Straattvtan, R. 1962. Notes on certain Lepidoptera ovipositing on plants which are toxic to their larvae. J. Lepid. Soc. 16: 99—103.
Verschaeffelt, E. 1910. The cause determining the selection of food in some herbivorous insects. Proc. Acad. Sci. Amsterdam 13: 536-542.
Voss, E. G. and W. H. Wagner. 1956. Notes on Pieris virginiensis .... hitherto unreported from Michigan. Lepid. News 10: 18-24.
THE BRAZILIAN "CERCYONIS" (SATYRIDAE)
Lee D. Miller
Allyn Museum of Entomology, 712 Sarasota Bank Building, Sarasota, Florida
and Thomas C. Emmel
Department of Zoology, University of Florida, Gainesville
Much confusion has occurred in the delimitation of the genus Cercyonis Scudder (1875). Periodically various authors have sought to unite these American butterflies with the Palearctic Satyrus Latreille (1810), and Minois Hiibner (1819) whereas other authors have pointed out the distinctness of Cercyonis. Miller (1968, pp. 99, 120) showed that the Nearctic Cercyonis are members of the satyrine tribe Maniolini, and in fact, the only American representatives of this basically Palearctic tribe,
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whereas Satyrus and Minois structurally belong to the Satyrini. The only representative of the Satyrini in the New World is Neominois Scudder (1875), from the western United States. Emmel (1969) has described the genus Cercyonis and by implication restricted it to the Nearctic.
The southern South American satyrid fauna has suffered from attempts to relate the butterflies found there to Holarctic genera that were familiar to the northern hemisphere systematists that first described the species. Many species belonging to the strictly American Pronophilini and Euptychini were described and long retained in such northern genera as Satyrus (Satyrini) and Epinephele (= Maniola: Maniolini). Such insects are illustrated particularly in the Elina and Lymonopodia series of the Pronophilini (Miller, 1968, pp. 117-118). A similar situation exists with regard to Cercyonis. Weymer (1912, pp. 228-230) listed the various Nearctic Cercyonis and includes the South American glaucope (C. & R. Felder) from southern Brazil and gustavi (Staudinger, 1897) from Bolivia. Forster (1964, p. 136) transferred the latter to the genus Argyr-ophorus Blanchard (1852), a pronophiline. The more recently described Cercyonis leuderivaldti Spitz (1931) must also be considered in this review of the extra-Nearctic "Cercyonis."
Should glaucope and leuderwaldti indeed be Cercyonis, a vast zoo-geographic problem would arise: how did these butterflies get from the Nearctic to the southern Neotropics without leaving intervening populations, when did this occur and by what route(s)? Furthermore, to which Nearctic species are these isolated populations related? The problem is purely academic, because leuderwaldti and glaucope are not even in the same tribe, and neither is a member of the Maniolini, as is Cercyonis.
For comparison the venation, palpus, male and female forelegs and the male genitalia of Cercyonis are given in Figs. 1-5.
Cercyeuptychia Miller and Emmel, new genus
Type-species: Cercyonis leuderivaldti Spitz. 1931. Rev. Erit. Sao Paulo 1: 46 (Brazil).
This genus is a member of the Euptychiini and conforms in the important characters with other members of the tribe, as defined by Miller (1968, pp. 90-92). A formal description follows:
Eyes naked. Antennae short, between two-fifths and one-half length of wing; club weakly developed, occupying distal quarter of antenna and slightly more than twice thickness of shaft at its thickest point. Palpi (Fig. 7) about two and a half times length of head, erect and slightly convergent at tips; th:;rd segment two-fifths length of second, hairs of second segment less than twice greatest segmental width.
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Journal of the Lepidopterists' Society
Figs. 1-5. Cercyonis pegala (Fabricius). 1, C. p. alope (Fabricius), $ venation (approx. 2x ), Connecticut, New Haven Co., Hamden (Allyn colln.); 2, same, palpus (approx. 12x) (LDM slide M-2107); 3, same, $ foreleg (approx. 12x ) (same slide as Fig. 2); 4, C. p. texana (Edwards), $ foreleg (approx. 12X ) (LDM slide M-2113); 5, C. p. alope, $ genitalia (approx. 12x ) (LDM slide M-2108).
Male foreleg (Fig. 8) reduced (forefemur-tibia-tarsus just over one-fifth length of same segments of midleg) with monomerous, unspined tarsus; femur somewhat longer than tibia. Female foreleg (Fig. 9) reduced, less than one-third length of midleg, femur somewhat longer than tibia, with a pentamerous, clubbed tarsus bearing spurs on the third and fourth subsegments. Ambulatory legs rather short, slender; midtibia less than twice length of proximal midtarsal subsegment, slightly spiny dorsad and with well-developed terminal spurs; midleg slightly shorter than hind leg.
Forewing cell square-out, slightly excavate along nii-m2, and about half length of forewing costa. Forewing radial veins arising from cell in two branches, Rs and Mi arising separately, M2 arising slightly near Mi than M:!, Cui arising nearer Ma than Cu2. Androconial patch of mealy and hairlike scales extending from 2A across cell to Mi-Ma, dentate distad in Ma-Ms. Sc and cubital stem inflated basally, 2A not. (Fig. 6).
Hind wing cell straight, slight distal migration of mo-m3 along My, and produced at origin of M3: length of cell to origin of M3 about three-fifths length of wing to end of M3. Vein 3A slightly longer than Sc Rx, M» and Ciu arising well separated and Ma arising nearer Mi than My. (Fig. 6).
Male genitalia (Fig. 10) typical of those of many euptychiines (Forster, 1964), especially as regards the free gnathos, a condition typical of many Euptychiini, but not in Maniolini or Pronophilini. Cenitalia similar to those displayed by genera
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Figs. 6-10. Cercyeuptychia leuderwaldti (Spitz). 6, $, venation (approx. 2x ) Brasil, D. F., Sobradinho, Brasilia (Allyn colln.); 7, palpus (approx. 12X ) (LDM slide M-2105); 8, $ foreleg (approx. 12x) (same slide as Fig. 7); 9, $ foreleg (approx. 12X) (LDM slide M-1724); 10, $ genitalia (approx. 12X ) (LDM slide M-2109).
Godartiana Forster (1964) and Fraefaunula Forster (1964), but spined gnathos typical of present genus.
The pattern of these butterflies (Figs. 16-19) is also reminiscent of Godartiana and Fraefaunula with the heavily striated under surface of both wings and the lack of distinct bands that are prominent in most other Euptychiini. The well-developed ocelli of Cercyeuptychia leuder-toaldti are only faintly indicated in any Godartiana, but well-developed in some Fraefaunula, and the wings of the present genus are rounded, as in Fraefaunula, not angular, as in Godartiana.
This remarkable genus is most closely related to Godartiana and Fraefaunula, but quite distinct from both and immediately recognizable by the spiny gnathos and much longer penis, and from Godartiana by the wing shape. One of us (LDM) is working currently on the Euptychiini and considers the present genus to be somewhat more advanced than either Godartiana or Fraefaunula, perhaps derived from one of them. K. S. Brown (in lift.) states that the present genus has comparable habits to Fraefaunula armilla (Butler).
The name of the genus is feminine and derived from the similarity of these butterflies to the Nearctic Cercyonis.
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Journal of the Lepidopterists' Society
Figs. 11—15. Pseudocercyonis glaucope (C. & R. Felder). 11, P. g. boenning-hauseni (Foetterle), $ venation (approx. 2x), Brasil, S. Paulo, Campos do Jordao (Allyn colln.); 12, same, palpus (approx. 12x ) (LDM slide M-2106); 13, same, $ foreleg (approx. 16x ) (same slide as Fig. 12); 14, P. g. glaucope (C. and R. Felder), 9 foreleg (approx. 16x ) (LDM slide M-1725); 15, P. g. boenninghauseni, $ genitalia (approx. 12X ) (LDM slide M-2110).
Pseudocercyonis Miller and Emmel, new genus
Type-species: Epinephele glaucope C. and R. Felder, 1867 [1864-1867]. Reise der . . . Fregatte "Novara" . . . Lep. Rhop., (3): 493-494; pi. 67, figs. 5, 6 (TL-"Brasilia").
This genus is a member of the tribe Pronophilini and conforms in major respects to the general characterization of that tribe by Miller (1968, pp. 110-114). A formal description follows:
Eyes naked. Antennae short, about two-fifths length of wing; club rather well developed, occupying distal quarter of antenna, about three times thickness of shaft and flattened at tip. Palpi (Fig. 12) about twice length of head, semi-porrect and somewhat divergent; third segment less than one-fourth length of second, hairs of second segment more than three times that of greatest segmental width.
Male foreleg (Fig. 13) greatly reduced (forefemur-tibia-tarsus only one-ninth length of those segments of midleg), with a stubby, monomerous, unspined tarsus; femur much longer than tibia. Female foreleg (Fig. 14) as aborted as that of male with a monomerous, unspined tarsus bearing no spurs; femur much longer than tibia. Ambulatory legs rather short and stubby; midtibia slightly more than twice
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length of proximal midtarsal subsegment, sparsely spiny dorsad and with well developed terminal spurs; mid- and hind legs subequal.
Forewing cell square-cut, slightly excavate along rrh-m2, and less than half length of forewing costa. Forewing radial veins arising from cell in two branches, Rs and Mi approximate but separate at their origins, M2 arising midway between Mi and M3 and Cui arising midway between M3 and Cu2. No distinct androconial patch. Sc greatly inflated at base, the cubital stem slightly inflated basad and 2A not at all (Fig. 11).
Hind wing cell "stepped" distad with a definite distal migration of m2-ma along M-> and produced at origin of M3: length of cell to origin M3 slightly less than half length of wing to end of M3. Sc-Ri subequal to 3A, Ms and Cui arising well separate and M2 arising somewhat nearer Ms than Mi (Fig. 11).
Male genitalia (Fig. 15) quite distinct from those of either Cercyonis (Fig. 5) or Cercyeuptychia (Fig. 10) but somewhat like those of Argyrophorus Blanchard, 1852 (Forster, 1964, pp. 135, figs. 168-169; Hayward, 1958, p. 254, fig. 43) and Tetraphlehia C, and R. Felder, 1867 (Hayward, 1958, p. 257, figs. 45, 47), but the longer, dorsally toothed penis immediately distinguishing the present butterflies.
The pattern of these butterflies (Figs. 20-21) is distinctive, but somewhat reminiscent of Tetraphlehia germainii C. and R. Felder, 1867 (Hayward, 1958, pi. 4, fig. 161). Only by stretching the imagination could glaucope be considered a Cercyonis, but it is not easily referred to Epi-nephele (— Maniola) in which it was described, either.
This singular genus resembles at least the type-species of Tetraphlehia superficially, but genitalically it is nearest Argyrophorus, the type of which is the amazing aluminum colored A. argenteus Blanchard (1852) from the mountains of Chile and Argentina. The venation of the present genus is rather close to that of Argyrophorus (Miller, 1968, p. 112, fig. 279), differing in minor details; the present genus does not have a distinct androconial patch. The elongate, dorsally toothed penis of this genus is characteristic. The present genus belongs to the Elina series of the Pronophilini (Miller, 1968, pp. 110, 117).
The generic name is feminine and refers to the fact that the type-species was wrongly considered a member of the Nearctic Cercyonis by many authors.
Discussion
The southern Neotropical "Cercyonis" are no more members of that Nearctic genus than is true Cercyonis synonymous with the Palearctic Minois. In fact, Cercyonis (Maniolini), Minois, (Satyrini), Cercyeuptychia (Euptychiini) and Pseudocercyonis (Pronophilini) are members of totally different tribes within the Satyrinae. The similar facies and the equivalent ecological niches shown by these four genera, as well as some South African Dirini, suggest a possible adaptive advantage to a morphological appearance such as shown by Cercyonis, etc., even though
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Journal of the Lepidopterists' Society
Figs. 16-19. Cercyeuptychia leuderwaldti (Spitz). 16, $ upper surface, Brasil, Goias, Ponte Funda (Emmel colln.); 17, same specimen, under surface; 18, $ upper surface, Brasil, Goias, Orizona (Emmel colln.); 19, same specimen, under surface.
Figs. 20, 21. Pseudocercyonis glaucope boenninghauserri (Foetterle). 20, £, upper surface, Brasil, S. Paulo, Campos do Jordao (Allyn colln.); 21, same specimen, under surface. Note: All figures approx. 1.5X-
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the butterflies are only remotely related. All appear to be scrub country species, some in low country, as some Cercyonis and Minois, others at high elevations, such as other Cercyonis and Pseudocercyonis. Such assemblages of ecological equivalents are not uncommon among the Satyridae, as demonstrated by the "Erebia series" of unrelated montane butterflies, including the Holarctic Erebia Dalman (1816) (Erebiini), the Lymanopoda series (Pronophilini) from the high Andes, Percno-daimon Butler (1876) and other New Zealand Hypocystini and some South African Dirini. Careful morphological examination is necessary on members of supposedly cosmopolitan, and particularly pantropical, groups to confirm or deny relationships that have all too long been taken for granted.
Acknowledgment
We should like to thank Dr. Keith S. Brown, Jr., of Bio de Janeiro, Brazil, for providing material and observations on the ecology of the south Brazilian "Cercyonis."
Bibliography
Emmel, T. C. 1969. Taxonomy, distribution and biology of the genus Cercyonis
(Satyridae). I. Characteristics of the genus. Jour. Lepid. Soc. 23: 165-175. Forster, W. 1964. Beitrage zur Kenntnis der Insectenfauna Boliviens XIX.
Lepidoptera III, Satyridae. Veroff. Zool. Staatssamml. Miinchen 8: 51-188. Hayward, K. J. 1958. Satiridos argentinos (Lep. Rhop. Satyridae) III. Guia
para su clasificacion. Acta. Zool. Lilloana 15: 199-295. Miller, L. D. 1968. The higher classification, phylogeny and zoogeography of
the Satyridae (Lepidoptera). Mem. American Ent. Soc. 24. Weymer, G. 1910-1912. Satyridae. in Seitz, A. Die Grossschmetterlinge der
Erde, vol. 5 (Die Amerikanische Tagfalter). Stuttgart.
CONSUL PANARISTE (NYMPHALIDAE) IN VENEZUELA
I secured two fresh males of Consul panariste (Hewitson) on 5 and 6 February 1968 while collecting in the Venezuelan Andes with Albert Gadou of Caracas. They were taken on banana bait at approximately 1000 meters elevation on the Barinitas to Santa Domingo road in the state of Barinas. This is a humid, tropical forest situation, transitional to cloud forest. Albert reported having taken previous examples of the species in the same location.
Comstock (1961. Butterflies of the American Tropics: The genus Anaea, p. 188) stated that Consul panariste is known to occur only in Colombia. C. panariste has been traditionally placed in the genus Anaea, before Comstock allied it with Consul fabius (Cramer) (= Protogonius hippona Fabricius). Although Comstock considered Consul to be a subgenus of Anaea, contemporary usage? usually elevates the subgenera in his monograph to generic rank.
John H. Masters, Lemon Street North, North Hudson, Wisconsin,